A simulation model is used to quantify relationships between diet quality, digestive processes and body weight in ungulate herbivores. Retention time of food in the digestive tract is shown by regression to scale with W, and to be longer in ruminants than in hindgut fermenters. Allometric relationships between whole gut mean retention time (MRT, h) and weight (W) were: MRT=9.4 W (r =0.80) for hindgut fermenters and MRT=15.3 W (r =0.76) in ruminants. Longer retention of ingesta by large-bodied ruminants and hindgut fermenters increases digestive efficiency relative to small animals and permits them to survive on lower-quality foods. Compared with ruminants, hindgut fermenters' faster throughput is an advantage which outweighs their lower digestive efficiency, particularly on poor quality foods, provided that food resources are not limiting. This suggests that the predominance of ruminants in the middle range of body weights results from their more efficient use of scarce resources under conditions of resource depletion. Considering only physical limitations on intake, the model shows that the allometric coefficient which scales energy intake to body mass is 0.88 in ruminants and 0.82 in hindgut fermenters. The advantages of large body size are countered by disadvantages where food quantity is limited, and we suggest that the upper limit to ungulate body size is determined by the ability to extract nutrients from feeding niches during the nadir of the seasonal cycle of resource quality and abundance.

The process of weaning is related to a critical or threshold body weight attained by offspring among large‐bodied mammals; the anthropoid primates, ungulates and pinnipeds. While weaning weight was allometrically related to maternal weight in interspecific comparisons, it was isometrically related to neonatal weight. When a neonate had grown to four times its birth weight, it was weaned. Differences between taxonomic groups were found only among the fasting phocids, where weanlings attained a lower, but proportional, weight. The duration of lactation was only weakly allometrically related to maternal or neonatal weight, and varied between individuals intraspecifically as a function of maternal condition. The time to weaning appears to be ecologically sensitive rather than to reflect interspecific life‐history variation, in that, irrespective of the time to weaning, similar proportional weights appear to be attained. Interspecific similarities in threshold weaning weights are suggested to result from constraints on maternal abilities to meet energetic requirements of offspring through lactation after infants attain a threshold weight.

Ruminants are unevenly distributed across the range of body sizes observed in herbivorous mammals; among extant East African species they predominate, in numbers and species richness, in the medium body sizes (10-600 kg). The small and the large species are all hind-gut fermenters. Some medium-sized hind-gut fermenters, equid perissodactyls, coexist with the grazing ruminants, principally bovid artiodactyls, in grassland ecosystems. These patterns have been explained by two complementary models based on differences between the digestive physiology of ruminants and hind-gut fermenters. The Demment and Van Soest (1985) model accounts for the absence of ruminants among the small and large species, while the Bell/Janis/Foose model accounts both for the predominance of ruminants, and their co-existence with equids among the medium-sized species (Bell 1971; Janis 1976; Foose 1982). The latter model assumes that the rumen is competitively superior to the hind-gut system on medium quality forages, and that hind-gut fermenters persist because of their ability to eat more, and thus to extract more nutrients per day from high fibre, low quality forages. Data presented here demonstrate that compared to similarly sized grazing ruminants (bovids), hind-gut fermenters (equids) have higher rates of food intake which more than compensate for their lesser ability to digest plant material. As a consequence equids extract more nutrients per day than bovids not only from low quality foods, but from the whole range of forages eaten by animals of this size. Neither of the current nutritional models, nor refinements of them satisfactorily explain the preponderance of the bovids among medium-sized ungulates; alternative hypotheses are presented.

A study was instigated to test whether grazing herbivores have evolved effective strategies to reduce parasite ingestion and to assess the effects of parasitism on these strategies. Two choice trials, each using five animals parasitized with O. circumcincta and five parasite naive animals, were conducted to investigate cues used by sheep to avoid grazing swards contaminated with faeces. Animals were presented with pairs of artificial swards (36 × 21 cm) and allowed to graze for short periods. In the first trial, four quantities (0 g, 5 g, 15 g and 25 g), and in the second trial a control (no faeces) and three age classes (0, 10 and 21 days old), of faeces were tested against each other. Increasing amount offaeces per sward tray was associated with a reduced proportion of bites taken from the sward and reduced bite depth and mass. The minimum amount of faeces at which sheep showed significant levels of rejection was 15 g. Decreasing age of faeces was associated with a reduced proportion of bites taken from the sward, reduced bite depth and mass. This presented the paradox that grazing swards contaminated with young faeces presented least risk of parasitism, yet fresh faeces presented the strongest stimulus for sward rejection. Parasitism altered animal grazing behaviour with parasitized animals becoming more selective when avoiding contaminated swards, taking smaller bites at reduced bite rates compared with non-infected animals. The enhancement of faeces avoidance behaviours shown by parasitized animals could act to reduce further intake of parasites and suggests that grazing behaviour is affected by nematode infection.

S U M M A R YSimulation modelling was used to investigate interactions between forage degradation characteristics, rumen processes and body weight, and to predict the voluntary food intake and digestion of a range of forages. Predicted voluntary intake and digestion agreed well with empirical data, explaining 61 and 70%, respectively, of variance in observed values. Since the data covered a wide range of animal weights and forage qualities, these results suggest that the model is a useful means of integrating the effects of animal and forage variables. Interactions were examined between animal weight and diet quality, as defined by the proportion of potentially digestible cell contents and cell walls and their rates of digestion. Retention time of food in the digestive tract was shown by regression to scale with W 027 . The time taken to comminute large fibre particles also scaled with W 027 . Longer retention of digesta by large ruminants increases digestive efficiency compared with small animals and would permit them to survive on lower-quality foods. The model showed that maximum intake of metabolizable energy scales with c. W 08 ', greater than the scaling of maintenance with W 073 . INTRODUCTIONThe principal functions of simulation modelling are to predict the behaviour and dynamics of complex systems, and to reveal where knowledge is lacking. Applied to comparative studies of nutritional physiology, a modelling approach has an additional benefit of allowing the results of empirical studies of a few species to be generalized to other species in the form of quantitative predictions. The strength of the comparative approach is that the underlying mechanisms can be used to establish general relationships across species, and this may throw into relief the deviations (assumed to be specific adaptations) of individual species from interspecific trends. In order to investigate these interspecific trends in the nutritional physiology of ruminants, a model of the relationships between ruminant body size, food characteristics, voluntary food intake and digestion was developed.The first aim was to develop a dynamic mechanistic model to simulate rumen processes and the fermentation characteristics of forages in order to predict voluntary intake and the extent of forage digestion.